Demographic Trends in The Cactaceae PDF

Demographic Trends in the Cactaceae
Author(s): Hctor Godnez-lvarez, Teresa Valverde, Pablo Ortega-Baes

Reviewed work(s):
Source: Botanical Review, Vol. 69, No. 2 (Apr. - Jun., 2003), pp. 173-203
Published by: Springer on behalf of New York Botanical Garden Press
Stable URL: http://www.jstor.org/stable/4354455 .
Accessed: 01/02/2012 19:26
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .
http://www.jstor.org/page/info/about/policies/terms.jsp
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of
content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms
of scholarship. For more information about JSTOR, please contact support@jstor.org.
New York Botanical Garden Press and Springer are collaborating with JSTOR to digitize, preserve and extend
access to Botanical Review.
http://www.jstor.org
The Botanical Review 69(2): 173-203
Demographic Trends in the Cactaceae

HECTOR GODiNEZ-ALVAREZ
Unidad de Biologia, Tecnologiay Prototipos
Facultad de Estudios SuperioresIztacala
UniversidadNacional Aut6nomade Mexico
Avenidade los Barrios 1, Los Reyes Iztacala
Tlalnepantla54090, Ap. Postal 314, Edo. de Mexico, Mexico
TERESA VALVERDE
Departamentode Ecologia y Recursos Naturales
Facultad de Ciencias
UniversidadNacional Autonomade Mexico
Ciudad Universitaria
Mexico City,D.F 04510, Mexico
AND
PABLO ORTEGA-BAES
Laboratorio de InvestigacionesBotanicas
Facultad de Ciencias Naturales
UniversidadNacional de Salta
Buenos Aires 177, Salta 4400, Argentina
I. Abstract/Resumen ...................................................
II. Introduction ...................................................
III. Distributionand AbundancePatterns ................
.....................
IV. Seed Germinationand Seedling Establishment
.
..................
V. Growthand Reproduction..............................................
VI. Size Structure...................................................
VII. PopulationDynamics ..................................................
A. ElasticityAnalyses .................................................
B. MatrixModels and Matrix Simulations.........
........................
VIII. Conclusions and Perspectives ...........................................
IX. Acknowledgments ...................................................
X. LiteratureCited ...................................................
XI. Appendix 1: DemographicInformation,Endangerment,and Geographical
Distributionof Cactus Species with Different Life-Forms .....
...........
Copies of this issue [69(2)] may be purchased from the NYBG Press,
The New York Botanical Garden, Bronx, NY 10458-5128, U.S.A.;
nybgpress@nybg.org. Please inquire as to prices.
Issued 16 December 2003

? 2003 TheNew YorkBotanicalGarden
173
174
175
177
178
183
187
189
191
194
194
195
195
202
174
THE BOTANICAL REVIEW
I. Abstract
Although our biological knowledge regardingcactus species is thoroughin many areas,
only in recentyears have ecologists addressedtheir demographicbehavior.Here we attempta
first review of the presentknowledge on cactus demography,includingan analysis of the published informationon species with differentgrowth forms and life-historytraits. Our review
shows that cactus distributionranges are determinedby environmentalheterogeneityand by
species-specific physiological requirements.Temperatureextremes may pose latitudinaland
altitudinaldistributionlimits. At a more local scale, soil propertiesdramaticallyaffect cactus
distribution.Most cacti show a clumpedspatialdistributionpattern,which may be the reflection of a patchy resourcedistributionwithin their heterogeneousenvironments.The association of cactiwith nurseplantsis anotherfactorthatmay accountforthis aggregateddistribution.
Many cacti grow in associationwith these perennialnurseplants,particularlyduringearlylifecycle phases. The shade provided by nurse plants results in reduced evapotranspiration
and
bufferedtemperatures,which enhancecactus germinationand establishment.In some cases a
certaindegree of specificity has been detected between particularcactus species and certain
nurse plants. Yet some globose cacti may establish in the absence of nurse plants. In these
cases, rocks and other soil irregularitiesmay facilitategerminationand establishment.
Cacti are slow-growing species. Several abiotic factors, such as water and nutrientavailability,may affect their growth rate. Competitionand positive associations(i.e., mycorrhizae
andnurse-cactiassociation)may also affect growthrate.Age at firstreproductionvariesgreatly
in relationto plantlongevity.In general,cactusreproductivecapacityincreaseswith plantsize.
Populationsare often composedof an unevennumberof individualsdistributedin the different
size categories. This type of populationstructurereflects massive but infrequentrecruitment
events, apparentlyassociatedwith benign periods of abundantrainfall.
A few cactusspecies have been analyzedthroughthe use of population-projection
matrices.
A total of 17 matriceswere compiled and compared.Most of them reflect populationsthatare
close to the numericalequilibrium(X = close to unity). Elasticity analyses revealed that the
persistenceof individualsin their currentsize category("stasis")is the demographicprocess
thatcontributesthe most to populationgrowthrate.Also, adultcategories(ratherthanjuveniles
or seedlings) show the largestcontributionsto X. No differenceswere apparentregardingthis
matterbetween cacti with differentlife-forms.This review shows thatourknowledgeof cactus
populationecology is still incipientand ratherunevenly distributed:some topics are well developed; for others the available informationis still very limited. Our ability to preservethe
greatnumberof cactus species thatarenow endangereddependson our capacityto deepenour
ecological understandingof their populationprocesses.
Resumen
A pesar de que nuestro conocimiento biol6gico sobre las cactaceas es basto en muchas
areas, ha sido s6lo hasta fechas recientes que los ec6logos han abordadoel estudio de su
comportamientodemografico.En este articulopresentamosuna primerarevisi6n del conocimientoactualsobredemografiade cactaceas,incluyendoun analisisde la informacionpublicada
sobreespecies con diferentesformasde crecimientoy caracteristicasde historiade vida.Nuestra
revisi6nmuestraque el areade distribuci6nde las cactaceasse ve afectadaporla heterogeneidad
del ambientey por los requerimientosfisiol6gicos de cada especie. Los valores extremos de
temperaturafijan limites latitudinalesy altitudinalesde distribuci6na muchascactaceas.A una
escala mas local, las propiedadesdel suelo juegan un papel fundamental.La mayoriade los
cactus muestranuna distribuci6nespacial agregada,lo cual es un reflejo de la distribuci6n
DEMOGRAPHY
IN THE CACTACEAE
175
aparchonadade los recursosen los ambientesaltamenteheterogeneosquehabitan.La asociaci6n

de los cactus con plantasnodrizaes otro de los factoresque explica su distribuci6nagregada.
Muchos cactus crecen en asociaci6n con estas plantas nodriza, particularmentedurantelos
primerosestadios.La sombrade estasplantasperennesreducela evapotranspiracion
y amortigua
la temperatura,lo cual incrementala germinaci6ny el establecimientode los cactus.En algunos
casos se ha detectadoespecificidadentreespecies particularesde cactusy ciertasplantasnodriza.
Aun asi, algunos cactos globosos pueden establecerse en ausencia de plantas nodrizas. En
estos casos, las rocas y otras irregularidadesdel terrenopodrianfacilitarla germinaci6ny el
establecimiento.
Los cactus son especies de lento crecimiento.Variosfactoresabi6ticos, como el agua y la
disponibilidadde nutrientes,pueden afectar su tasa de crecimiento. La competencia y las
asociacionespositivas (i.e., formacionde micorrizasy asociacioncon plantasnodriza)tambien
pueden afectar su tasa de crecimiento.La edad a la primerareproduccionvaria mucho con
relaci6n a la longevidad de las plantas. En general, la capacidadreproductivade los cactus
Las poblacionesfrecuentementeestaincompuestasde
aumentaconformeaumentasu tamainio.
individuosdistribuidosde manerairregularentrelas diferentesclases de tamanlo.Este tipo de
estructura poblacional refleja eventos masivos pero poco frecuentes de reclutamiento,
aparentementeasociados a periodosbenignos de abundanteslluvias.
Solo unas cuantasespecies de cactus se han estudiadoa traves de matricesde proyecci6n
poblacional.En este estudio se compilarony se compararon17 matrices.La mayoriade ellas
reflejanpoblacionesque se encuentrancerca del equilibrionumerico(X = cerca de la unidad).
Los analisisde elasticidadrevelaronque la persistenciade los individuosen su mismacategoria
de tamafio ("estasis") es el proceso demografico que mayormentecontribuye a la tasa de
crecimientopoblacional.Tambiense vio que las categoriasde adultos(y no las de juveniles o
plantulas)fueronlas que mostraronun mayoraportea X.No se encontrarondiferenciasa este
respectoentreespecies de cactuscon diferenteformade crecimiento.Estarevisi6nmuestraque
nuestro conocimiento sobre la ecologia poblacional de los cactus es aun incipiente y esta
distribuidode manera dispareja:algunos temas estan bien desarrolladosmientrasque para
otrosla informaci6ndisponiblees aunmuy limitada.La posibilidadde conservarel grannuimero
de especies de cactaiceasque se encuentranamenazadasdepende de nuestracapacidadpara
profundizaren el entendimientoecol6gico de sus procesos poblacionales.
II. Introduction
Cactiare a very diverseandcomplexplantfamily comprisingmorethan2000 species, most
of whicharenativeto aridandsemi-aridenvironmentsin theAmericas(Bravo-Hollis& SanchezMejorada,1978, 1991). The extraordinaryadaptationsof these plantsto low wateravailability
and other stressfulconditionshave made them the subjectof a numberof morphologicaland
physiological studies, includingaspects relatedto theirphotosyntheticpathways,their waterstorage ability, and the various other ways in which they optimize their water-useefficiency
(amongothers,Bravo-Hollis& Sanchez-Mejorada,1978, 1991;Gibson& Nobel, 1986;Nobel,
1988; Altesor et al., 1992). This information,along with a host of botanicalrecordsand taxonomic analyses, has been tremendouslyvaluable in aiding our understandingof the biology
and evolutionof this fascinatingplantfamily.Yet the lack of ecological informationon cacti is
still notoriousin the biological literature.In recentyears a numberof authorshave published
ecological studieson cacti, coveringaspect such as reproductivebiology (Tinoco-Ojanguren&
Molina-Freaner,2000; Fleming et al., 2001), plant-pollinatorand plant-disperserinteractions
(Fleming & Valiente-Banuet,2002; Godinez-Alvarezet al., 2002), the ecophysiology of ger-
176
minationandearlygrowth(Rojas-Arechiga& Vazquez-Yanes,2000; Ruedaset al., 2000; Flores

& Briones,2001; Rojas-Arechigaet al., 2001), and cactus-nurseplantinteractions(McAuliffe,
1984a, 1984b;Valiente-Banuet& Ezcurra,1991; Valiente-Banuetet al., 1991a, 1991b; FloresMartinezet al., 1994, 1998). Additionally,recent articleshave addressedthe demographyof
cactus populations,including species with contrastingdistributionranges, growth forms, and
life-history traits. It is this kind of ecological informationthat we are concernedwith in the
present article. We believe that our present knowledge of the demographyof cactus species
deservesa first attemptto searchfor patternsandtrendsthatmay influencethe futuredevelopment of this particulararea of plant ecology. A detailedevaluationof the presentstatusof our
knowledge will allow us to establish priorities and encourage researchersto develop ideas
aboutparticularissues that may eventuallyfill the gaps on this subject.
Partof the need to studythe demographictrendsemergingin the Cactaceaeis relatedto the
fact thatthere is a high numberof threatenedspecies in this plant family (Anderson& Taylor,
1994; Hunt, 1999). In fact, the whole family is includedin appendix2 of the CITESbook of
endangeredspecies (Hunt, 1999). Some of the reasonsthat may accountfor this are: 1) Many
cacti are highly restrictedin their distributionand occupy very specific habitats,which makes
them prone to extinction by habitatdestructionand land-use change (Esparza-Olguinet al.,
2002); 2) They seem to be particularlysensitive to disturbancesdue to their low individual
growthratesandhighlyvulnerableearlystagesof development(Hermnndez
& Godinez-Alvarez,
1994); 3) Illegal collection and tradeare an increasingpressureon many cactus populations,
especially since they have become more fashionablein recentyears; and 4) Theirhabitatsare
frequentlyassociatedwith poor areasin developingcountries,where the pressureof increasing
human populationson land-use change toward farming and cattle ranchingare very strong.
Thus the increasingdemographicinformationon cactus species will certainlyaid in the design
of conservationand managementplans and will offer a tool with which to evaluatethe conservation statusof poorly known species in this plant family.
In additionto conservationand managementinterests,demographicinformationis the basis on which our understandingof populationdynamics and life-history traits is supported
(Silvertownet al., 1993). Recent demographicinformationon cactus species is addingsignificantly to our understandingof plant populationdynamics in nature,since they include longlived species that inhabitdry tropicalareas, two particularitiesthat are poorly representedin
the demographicliterature.In this articlewe attempta firstreview of the demographicfeatures
of cactus species, accordingto the literaturepublishedto date,which is summarizedin Appendix 1. A decade ago Silvertown et al. (1993) published their first literaturereview on plant
demographictrends.In thatreview, in which interestingtrendsbetween demographyand lifeform emerged, only one cactus, Carnegiea gigantea, was included. In this article we use the
approachdevelopedby Silvertownet al. (1993) to analyzethe informationon cactusdemography publishedmainly in the last decade.We use a comparativeapproachthatfocuses on cactus
species with differentlife-forms, since a correlationbetween life-formand some demographic
propertiesappearsto be emergingamong desertplants (Flores & Briones, 2001). We consider
the following life-forms,accordingto the classificationused by otherauthors(Gibson& Nobel,
1986):columnarcacti(specieswith long, column-like,ribbedstemsmeasuring>2.0 m in height);
barrelcacti (plantswith ribbedstems, with a maximumheight of 0.5-2 m); globose cacti (species with single or multiple hemisphericalstems); cylindropuntias(opuntiaswith cylindrical
stems); and platyopuntias(opuntiaswith flattenedstems). In additionto data on population
dynamics,we includevaluableinformationon otheraspectsof cactuspopulationecology, such
as distributionand abundancepatterns,data on growth, reproduction,and populationstructure, and survival and establishmentduring early life-cycle phases. Our aim is to search for
DEMOGRAPHY
IN THE CACTACEAE
177
trendsandpatternsthatmay contributeto ourunderstandingof the factorsthatdeterminelongterm populationdynamics in nature,which still remaina centralissue in ecology (Horvitz&
Schemske, 1995).
III. Distribution and Abundance Patterns

Studieson the distributionandabundancepatternsof cacti have shownthatthese ecological
attributesmay vary both temporallyand spatially(Bowers et al., 1995; de Viana, 1996-1997;
Bowers, 1997b;Mourelle& Ezcurra,1997; Parker,1988a, 1993). Withregardto the distribution of cacti at a regional scale, environmentalheterogeneityas well as the particularphysiological requirementsof each species appearto determinetheirdistributionrange.Forinstance,
the columnarcacti Carnegiea gigantea, Lophocereusschottii, and Stenocereusthurberioccupy different habitats along the topographicand edaphic gradientsof the Sonoran Desert
(Parker,1988a). In addition,the northerndistributionlimits of these species appearto be determinedby the occurrenceof low temperatures,which severelydamagetheirstem apices (Nobel,
1980). Mourelleand Ezcurra(1997) also mentionthat variationin environmentalfactorssuch
as temperature,topography,and rainfallmay be associated with the occurrenceof different
Argentinecacti.
At a local scale, the three spatialdistributionpatternsdescribedin the ecological literature
(random,uniform,and clumped)have been found in cactus populations(Gulmonet al., 1979;
de Viana et al., 1990; Martinezet al., 1993; Huerta& Escobar, 1998; Valverdeet al., 1999;
Cody,2000). Yet clumpeddistributionsappearto be farmore commonthanrandomor uniform
distributions.It has been suggestedthat the aggregationof individualsthat has been found in
cacti with differentlife-forms(i.e., Capiapoacinerea,Ferocactushystrix,Ariocarpustrigonus,
Mammillariagaumeri, M magnimamma,M oteroi, Turbinicarpus
pseudopectinatus,Trichocereuspasacana, and Neobuxbaumiamacrocephala)may be relatedto the patchydistribution
of conditions that enhance seedling establishmentand plant growth (de Viana et al., 1990;
Valverdeet al., 1999; Martinezet al., 2001; Esparza-Olguinet al., 2002), which is bound to
exert a particularlystrongeffect on those species thatrequirethe shadeof a nurseplant.Other
potentialmechanismsthat may also account for a clumped distributionpatternare restricted
seed dispersal(Martinezet al., 1993, 1994) and patchy distributionof seeds in the soil (Hutto
et al., 1986; de Viana, 1996-1997; de Viana et al., 2001). Additionally,some studies have
shown thatyoung andadultplantswithin a populationmay have differentdistributionpatterns.
Intraspecificinteractions,such as competitionleading to self-thinningprocesses, may be responsible for this variation(de Vianaet al., 1990).
In regardto abundancepatterns,it is difficultto find a generaltrendamong cactus species
because populationdensities may vary greatly,from very few individualsper hectareto thousands per hectare(Valiente-Banuet& Ezcurra,1991; Martinezet al., 1993, 1994; Schmalzel
et al., 1995; de Viana,1996-1997;Huerta& Escobar,1998;Mandujanoet al., 1998;Valverdeet
al., 1999;Esparza-Olguin
et al., 2002). The density of a populationis the result of its birthand
deathrates, which may be affected by a host of biotic and biotic factorsat regional and local
scales. Apparently,massive but infrequentrecruitmentevents are associated with favorable
rainfall and temperatureperiods. In particular,infrequentevents such as "El Ninfo"may increasethe numberof new recruitsin some populations.This has been suggestedfor the columnarcacti Carnegieagigantea andStenocereusthurberi(Parker,1993; Pierson& Turner,1998),
the barrelcacti Echinocactuspolycephalus,Ferocactus acanthodes,and F. cylindraceus(Jordan & Nobel, 1981; Goldberg& Turner,1986; Bowers et al., 1995; Bowers, 1997b), and the
platyopuntiasOpuntia acanthocarpa, 0. basilaris, and 0. erinacea (Bowers et al., 1995).
178
Contrastingly,Haman(2001) reportsthatEl Nino events have an adverseeffect on two species

of Opuntia(0. echios and 0. galapageia) in the GalapagosIslands, by increasingseedling,
juvenile, and adultmortalitydue to waterlogging following heavy rainfall.In additionto rainfall patterns,otherfactorssuch as the length of the dry period,the availabilityof safe sites for
establishmentand growth,the presenceof trees and shrubsin the habitat,soil erosion, andthe
effect of livestockgrazingareimportantin determiningcactusdensityat a particularsite (Turner
et al., 1966; Jordan& Nobel, 1981, 1982; Nobel, 1989; Valiente-Banuet& Ezcurra, 1991;
Parker,1993; Bowers et al., 1995; Bowers, 1997b;Pierson& Turner,1998; Esparza-Olguinet
al., 2002).
Soil characteristicsappearto play an importantrole in the distributionand abundanceof
cactusspecies in aridandsemi-aridlandscapes(Parker,1991). The edaphicconditionsin which
some species live may be highly specialized.It has been observedthat columnarcacti such as
Cephalocereuscolumna-trajani,Escontriachiotilla,Neobuxbaumiatetetzo,Pachycereusfulviceps, P pringlei, Stenocereusgummosus, and S. thurberi,as well as globose cacti such as
Mammillariacrucigera,areassociatedwithparticularsoil types (Meyran,1980;Valiente-Banuet
et al., 1995; Ortega-Baes,2001; Contreras& Valverde,2002; Valiente-Banuet& GodinezAlvarez, 2002). Certainedaphicproperties,such as soil texture,may dramaticallyaffect water
availability,thus preventingthe establishmentof certainspecies while favoringothers,depending on theirparticularwaterrequirementsand tolerancesfor germinationand early growth.
IV. Seed Germination and Seedling Establishment

A greatnumberof studieshave addressedthe germinationbehaviorof cactusspecies under
both experimentaland naturalconditions. Since Rojas-Arechigaand Vazquez-Yanes(2000)
recently publisheda thoroughreview of the germinationecology of cacti, in this section only
some generalaspects will be discussed.
The main factorsthat affect seed germinationare water,temperature,and light. However,
embryoimmaturity,salinity,seed age, planthormones,inhibitorycompoundsin the testa,plant
domestication,and other factors may also affect this process (Rojas-Arechiga& VazquezYanes,2000; Rojas-Arechigaet al., 2001). Of all these factors,wateravailabilityappearsto be
the most importantfor germinationof cactusseeds, since it is the most limitingfactorunderthe
conditions that prevail in deserts. Dubrovsky(1996, 1998) found that seeds of the columnar
cacti Carnegieagigantea, Pachycereuspecten-aboriginum,Stenocereusgummosus,andS. thurberi, as well as seeds of the barrelcactusFerocactuspeninsulae, germinatedmore rapidlyand
accumulatedhigher biomass when they were subjectedto hydration-dehydration
cycles. This
phenomenonis known as "seedhydrationmemory"andmay facilitatethe germinationprocess,
leading to an increasein the probabilitythat seedlings will survive (Dubrovsky,1996).
Most studies of seed germinationof cactus species have been conductedundercontrolled
conditions, and only a few have consideredtheir germinationbehavior in the field. Further
field researchis necessary in this area, since there are importantaspects of seed ecology of
which the demographicimplicationsare ignored.One of these aspectsis the potentialof cactus
seeds to remainviable in the soil for long periodsof time, forminglong-termburiedseed banks
(Rojas-Ar6chiga& Vazquez-Yanes,2000; Rojas-Ar6chiga& Batis, 2001). The only available
informationon this subject suggests that seeds of the barrelcactus Ferocactus wislizeni may
remainviable in the soil for at least 18 months;postdispersalseed predationin this species is
high, althoughseeds may escape from predatorswhen hidden among rocks (Bowers, 2000).
Also, Ortega-Baes(2001) foundviable seeds of Escontriachiotilla of unknownage (at least 12
monthsold) buriedin the soil, and field experimentswith Mammillariamagnimammashowed
DEMOGRAPHY
IN THE CACTACEAE
179
that its seeds may remain viable in the soil for more than a year (Valverdeet al., in press).
Additionally,several Mammillariaspecies have been reportedto retain fruits in their stems,
which may play a similarecological role as a buriedseed bank (Rodriguez-Ortega& Franco,
2001).
Studies of seed germinationand seedling survival of cactus species under naturalconditions have shown that the presence of perennialplants is necessary for the successful recruitment of new individuals(Niering et al., 1963; Tumeret al., 1966; Steenbergh& Lowe, 1969,
1977; Jordan& Nobel, 1981, 1982; McAuliffe, 1984a; Valiente-Banuet& Ezcurra, 1991;
Mandujanoet al., 1998). Under the harsh environmentalconditions that prevail in deserts,
these perennialnurseplantsfunctionas islandswithin which physical and/orbiotic conditions
are beneficial for the establishmentand growth of young cacti (Callaway, 1995). The nurse
plantsdecreasethe contrastbetweenmaximumandminimumsoil temperatures(Shreve, 1931;
Jordan& Nobel, 1981; Hutto et al., 1986; Valiente-Banuetet al., 1991a;Arriagaet al., 1993;
Suzan et al., 1996), provideprotectionagainstpredators(Steenbergh& Lowe, 1977; Huttoet
al., 1986;Valiente-Banuet& Ezcurra,1991), increasesoil-nutrientavailabilityundertheircanopies (Garcia-Moya& McKell, 1970; Nobel, 1989), andprovidea shadyenvironmentin which
evapotranspirationis substantiallyreduced (Tumer et al., 1966; Steenbergh& Lowe, 1977;
Nolasco et al., 1997; Godinez-Alvarez& Valiente-Banuet,1998). Additionally,they constitute
roostingsites for dispersers,therebyattractingimportantamountsof seed dispersal(Steenbergh
& Lowe, 1977; Yeaton, 1978; Hutto et al., 1986; Godinez-Alvarezet al., 2002).
The literatureon the nurse-plantphenomenonin cacti is summarizedin Table I. The first
point we may note from this table is that the associationwith perennialplants occurs in all of
the life-forms that are representedin the cactus family. For instance, Suzainet al. (1996) reportedthat 17 cactus species from the SonoranDesert with differentlife-formsgrow in association with the long-lived tree Olneya tesota. Also, Arriagaet al. (1993) reportedthat the
numberof individualsof Stenocereusthurberiand Ferocactuspeninsulae was significantly
higher underthe canopy of perennialplants than in open spaces in the tropicaldry forests of
northwestemMexico. In the TehuacanValley of centralMexico, five cactus species with differentlife-forms were found to be associatedwith differentperennialplants (Valiente-Banuet
et al., 199la). In SouthAmerica,the columnarcactus Trichocereusatacamensisshows a positive associationwith trees and shrubs(de Viana, 1996-1997; de Vianaet al., 2001). In most of
these cases it has been suggestedthatthe shadeprovidedby nurseplantis the main factorthat
enhancesseedling establishmentby bufferingextremetemperaturesand reducingevapotranspiration.
The identityof nurseplants so far reportedis varied (TableI). Differentplant species may
play this role, includingshrubs,trees,grasses,agaves, andeven cacti (McAuliffe, 1984b;Hutto
et al., 1986; Nobel, 1989; Valiente-Banuetet al., 1991a; Suzan et al., 1996; Mandujanoet al.,
1998; de Vianaet al., 2001; Ortega-Baes,2001). The numberof cactus seedlings andjuveniles
establishedbeneaththe canopiesof some nurse-plantspecies is higherthanexpectedby chance,
accordingto theirrelative abundancein theirplant communities(Huttoet al., 1986; ValienteBanuetet al., 1991a, 1991b;Mandujanoet al., 1998; de Vianaet al., 2001; Ortega-Baes,2001).
This pattem suggests that, in some cases, the nurse-cactus association may have reached a
certaindegree of specificity,which may be relatedto particulartraits(i.e., morphology,plant
architecture,phenology) that determinedifferences in the quality of each perennialplant as
nurse for different cactus species (Callaway, 1998). Certaintraits, such as the presence of
spines,deeproots,long-livedleaves, secondarymetabolites,andnitrogen-fixationability,among
others,may favor the establishmentof new individualsunderthe canopy of specific perennial
plants (McAuliffe, 1984a; Suzan et al., 1996; Callaway,1998).
Table I
=
Association between nurse plants and different cacti species

species with a higher number of recruits growing in association with nurse plants than expected
A= adults; Mechanism = advantages offered by the nurse plant to the growing cacti; ? = un
Species
Columnarcacti
Carnegiea gigantea*
Nurse plant
Ambrosiadeltoidea
Cercidiummicrophylum
Enceliafarinosa
Larrea tridentata
O/neya tesota
Prosopisjuliflora
Cephalocereus hoppenstedtii* Caesalpinia melanadenia
Escontria chiotilla*
Acacia cochliacantha
Fouquieriaformosa
Mimosa luisana
M. polyantha
Zizyphuspedunculata
Lophocereus schottii*
Celtis pallida
Lycium andersonii
L. berlandieri
Olneya tesota
Zizyphusobtusifolia
Neobuxbaumiamacrocephala Lippia graveolens?
N. tetetzo*
Caesalpinia melanadenia
Castela tortuosa
Eupatoriumodoratum
Mimosa luisana
Pachycereus pringlei
O/neya tesota
Prosopis articulata
Stenocereus thurberi
Haematoxylon brasiletto
Jatropha vernicosa
Tecomastans
Life-cycle stage Mechanism
S-J-A
Shade
Frost protection
Predationavoidance
S-J
S-J-A
Shade
Shade
S-J
Shade
Frost protection
S-J?
S-J
Shade?
Shade
S-J
Shade
Increased soil nutrients
Shade
S-J-A
Trichocereusatacamensis*
Aphyllocladus spartoides
Larrea divaricata
Prosopisferox
Barrel cacti
Echinocereus conglomeratus
Euphorbia antisyphilitica
Hamatocactus hamatacanthus
Opuntia leptocaulis
Opuntiafulgida
E. engelmannii*
Echinomastus erectrocentrus Celtis pallida
Lyciumandersonii
L. berlandieri
Olneya tesota
Zizyphusobtusifolia
Ambrosiadumosa
Ferocactus acanthodes
Ephedra aspera
Hilaria rigida
Hilaria rigida
F. cylindraceus
Haematoxylon brasiletto
F. peninsulae
Hamatocactus hamatacanthus Euphorbia antisyphilitica
Opuntia leptocaulis
Celtis pallida
Peniocereus greggii
Lyciumandersonii
P. striatus
L. berlandieri
Olneya tesota
Zizyphusobtusifolia
Globose cacti
Ariocarpus trigonus
Coryphanthapallida*
Mammillaria casoi*
M carnea
M colina*
Castela tortuosa
Eupatoriumodoratum
Caesalpinia melanadenia
Castela tortuosa
Castela tortuosa
Eupatoriumodoratum
S-J-A
S-J-A
S-J-A
Shade
Predation avoidance
Shade
Shade
S-J-A
S-J-A

Shade
S-J-A
S-J-A
Shade
Shade
S-J-A
S-J-A
Shade?
Shade
S-J-A
-
S-J-A
Shade
-
Shade
Table I, continued
Species
Globose cacti, continued
M. gaumeri
M. microcarpa
M oteroi
M. thornberi
Cylindropuntias
Opuntiaacanthocarpa
0. echinocarpa
0. leptocaulis
0. ramosissima
Platyopuntias
0. rastrera*
0. streptacantha
Nurse plant
Life-cycle stage Mechanism
S-J
Opuntiafulgida
Quercus castanea
Celtispallida
Lyciumandersonii
L. berlandieri
Olneya tesota
Zizyphusobtusifolia
Hilaria rigida
Krameria sp.
Hilaria rigida
Krameria sp.
Larrea tridentata
Hilaria rigida
Krameria sp.
Cordiaparvifolia
Hilaria mutica
Jatrpoha dioica
Prosopis glandulosa
Acacia schaffneri
S-J-A
Shade?
-
Shade
S-J
-
S-J
S-J-A
Seed dispersal site?

-
Shade
Predation avoidance
Shade
IN THE CACTACEAE
DEMOGRAPHY
183
Despite the fact thatnurseplantsmay offer cacti some protectionduringthe initiallife-cycle

phases, mortalityis tremendouslyhigh at this stage. Survivorshipcurves of cactus seedlings
growingunderthe canopiesof differenttrees and shrubsaremainlytype III (sensu Pearl, 1928;
see Appendix 1; Tumeret al., 1966; Steenbergh& Lowe, 1969; Valiente-Banuet& Ezcurra,
1991; Mandujanoet al., 1998; Esparza-Olguinet al., 2002; Valverdeet al., in press). Yet the
intensityof seedlingmortalityis lowerbeneaththe canopiesof nurseplantsthanin open spaces.
For instance, seedlings of Mammillariagaumeri associatedwith nurse plants showed a constant mortalityrate throughtime (i.e., type II), whereas those growing in bare soil showed a
clear type III curve (Leirana-Alcocer& Parra-Tabla,1999).
Althoughthe evidence discussedso far may suggest thatthe nurseplant-cactusassociation
is prevalentthroughoutthe Cactaceaefamily (Table I), we must bear in mind that there are a
number of examples in which new cactus seedlings have been reportedto establish in the
absenceof nurseplants. Curiously,these species are mainly globose cacti, such as Ariocarpus
fissuratus,Epithelanthabokei,Mammillariamagnimamma,M lasiacantha,M pectinifera,and
Turbinicarpus
pseudopectinatus,which may grow in open spaces(Nobel et al., 1986;Arriagaet
al., 1993; Martinezet al., 1994;Valverdeet al., 1999; Rodriguez-Ortega& Ezcurra,2001). For
otherdesertplantsthe presenceof rocks on the ground,as well as otherirregularities,appearto
be of vital importancefor seed germinationand/orseedling survival.Rocks may reduce solar
radiationand thermalloadings,as well as prolongthe presenceof soil moisturein their immediate vicinity (Larmuth& Harvey, 1978). Thus the importanceof rocks and other surface irregularities(cavities, holes, etc.) as potentialfacilitatorsof germinationand seedling survival
in differentcactus species is one of the areasto which furtherresearchshouldbe directed.
V. Growth and Reproduction

After seed germinationand seedling establishment,growth and reproductionof cacti may
be affectedby extrinsicfactorssuch as biological interactionsandenvironmentalconditions,as
well as by intrinsicfactorssuch as genetic variation,allometry,andmetabolicrates.Biological
interactionswith other plants, animals, or microorganismsmay have a positive or negative
impact on cacti. During early growth, positive interactionssuch as the association with vesicular-arbuscularmycorrhizalfungi can enhance biomass productionby increasingnutrient
absorptionefficiency. This appearsto be the case of the columnarcactusPachycereuspectenaboriginum in a tropicaldry forest in Mexico (Rincon et al., 1993). The presence of mycorrhizae has also been reportedin Mammillariamagnimamma(Ruedas et al., 2000), although
theireffect on the growth of this species has not been analyzed.Otherbiotic interactionsmay
negatively affect the growth of cacti; such is the case of intra-and interspecificcompetition.
Paradoxically,these interactionsare enhancedunder the canopies of trees and shrubswhere
seedling establishmentis most successful.McAuliffe andJanzen(1986) documentedthe effect
of intraspecificcompetitionfor limitedwaterin denseaggregationsof young Carnegieagigantea
plants associatedwith the canopy of perennialshrubs;in this case, the effects of competition
were decreases in water uptake, water storage, apical growth, and reproductivepotential
(McAuliffe & Janzen, 1986). It has also been reportedthat some columnarcacti (C. gigantea
andNeobuxbaumiatetetzo)may establishintenseinterspecificcompetitionfor waterwith their
nurse plants, producinga decline in leaf, flower, and fruit productionin the nurse trees and
shrubs (McAuliffe, 1984a; Flores-Martinezet al., 1994, 1998). This type of interactionhas
also been documentedbetween the barrel cactus Ferocactus acanthodes and the perennial
bunchgrassHilaria rigida (Franco& Nobel, 1989), between Opuntialeptocaulisandthe shrub
Larreatridentata(Yeaton,1978), andbetween 0. rastrera,the perennialtussock grassHilaria
mutica,and the shrubL. tridentata(Briones et al., 1998). Furtherresearchis needed to deepen
184
THE BOTANICALREVIEW
our understandingof the way in which both positive and negative interactionsaffect cactus
growth.
In additionto biotic interactions,cactus growth may be affected by environmentalconditions. The literatureon this subject indicates that winter precipitationand frost decrease the
growthrateof Lophocereusschottii andStenocereusthurberi(Parker,1988b). In contrast,high
summerand low wintertemperatures,soil-moistureavailability,and pre- and postsummerdry
periods are among the main environmentalfactors that affect the growth rate of Carnegiea
gigantea (Steenbergh& Lowe, 1977). Experimentalanalyses of the early growthof cacti also
show that it is affected by environmentalconditions. For instance, a high concentrationof
nutrientssignificantlyincreasedseedling growth in Mammillariamagnimamma,Pachycereus
hollianus, and P. pringlei (Godinez-Alvarez& Valiente-Banuet,1998; Carrillo-Garciaet al.,
2000; Ruedaset al., 2000). Effects of directsolarradiationwere contradictory,for it increased
seedling growth in M. magnimamma(Ruedas et al., 2000) but decreased the growth of
Neobuxbaumiatetetzo, Opuntiarastrera,P hollianus, P pringlei, and S. thurberiseedlings
(Nolasco et al., 1997; Godinez-Alvarez& Valiente-Banuet,1998; Mandujanoet al., 1998;
Carrillo-Garciaet al., 2000).
Genetic variationamong individuals,as well as allometry,are also among the factorsthat
affect growthrates.Withregardto allometry,there is some variationin the annualgrowthrate
of Carnegieagigantea relatedto changes in plant form associatedwith ontogeny (Hastings&
Alcorn, 1961; Steenbergh& Lowe, 1977). Thus,afterseedling establishment,individualsgrow
rapidlyuntilthey reacha height of about2 m. This fast growthis the resultof an increasein the
photosyntheticsurface area and water-storagecapacity of individuals.However, growth rate
decreaseswhen plantsarebetween2 and6 m tall, since they startusing some of the productsof
photosynthesisto generatereproductivestructures.In largerplantsgrowthoccursmainlythrough
the proliferationof branches,while apparentlythe mainstemreachesa maximumheight(Shreve,
1910; Hastings & Alcom, 1961; Steenbergh& Lowe, 1977).
The growth patternis differentfor other species of columnarcacti, such as Lophocereus
schottii and Stenocereusthurberi.In these species the annualgrowth rate increases with increasing plant size (Parker,1988b). Differences in growth rate among species of columnar
cacti may be relatedto the particularbranchingpatternof each species. For instance,L. schottii
and S. thurberigrow throughthe productionof new stems, whereas Carnegieagigantea produces a single stem with severalbranches.
Since all cactusspecies are slow-growing,long-livedplants,thereis little informationabout
the relationshipbetween size and age (Table II),and thereforeage-relatedlife-historytraits
have been difficult to investigate.The availableinformationshows that age at first reproduction is variable, depending on the species, life-form, and plant longevity. In the long-lived
columnarcacti,plantsreproduceforthe firsttime when individualsare33 yearsold in Carnegiea
gigantea (Steenbergh& Lowe, 1977), 70 years old in Cephalocereuscolumna-trajani(ZavalaHurtado& Diaz-Solis, 1995), and more than 90 years old in Neobuxbaumiamacrocephala
(Esparza-Olguinet al., 2002). For a shorter-livedglobose cactus (Mammillariamagnimamma)
and a platyopuntia(Opuntia engelmannii),first reproductionoccurs when individualsare 4
and 9-11 years old, respectively(Bowers, 1996a;Valverdeet al., in press).
Once plantshave reachedreproductivesize, fecundityappearsto increasewith increasing
size (Fig. 1; Appendix 1). This patternwas observedin species with differentlife-forms, such
as the globose cacti Coryphantharobbinsorum,Mammillariacrucigera, M magnimamma
(Schmalzelet al., 1995; Contreras& Valverde,2002; Valverdeet al., in press),the platyopuntia
0. rastrera(Mandujanoet al., 2001), the barrelcactusEchinomastuserectrocentrus(Johnson,
1992; Johnsonet al., 1992), and the columnarcacti Carnegieagigantea, Escontria chiotilla,
DEMOGRAPHYIN THE CACTACEAE
185
Table II
Age or size at which first reproduction occurs in cacti with different life-forms
Age
(years)
Size
(meters)
Columnarcacti
Carnegiea gigantea
Cephalocereuscolumna-trajani
Lophocereusschottii
Stenocereus thurberi
33
70
-
2.2
3.3
>2.0
2.0-2.5
4-l oa
Barrel cacti
Echinomastuserectrocentrus
0.024
Johnson et al., 1992
Globose cacti
Coryphantharobbinsorum
Mammillariamagnimamma
0.013
-
Schmalzel et al., 1995

Valverdeet al., in press
9-11
6-13b
Bowers, 1996a
Species
Opuntioidcacti
Opuntiaenglemannii
Source
Steenbergh& Lowe, 1977
Zavala-Hurtado& Diaz-Solis, 1995
Parker,1989
Parker,1987
Plant size measuredas the numberof branches.

bPlant size measuredas the numberof cladodes.
Lophocereus schottii, Neobuxbaumia macrocephala, N. tetetzo, and Stenocereus thurberi

(Steenbergh & Lowe, 1977; Parker, 1987, 1989; Godinez-Alvarez et al., 1999; Ortega-Baes,
2001; Esparza-Olguin et al., 2002). It has been suggested that this pattem is related to an
increase in the number of branches as plants age, at least in columnar cacti (Steenbergh &
Lowe, 1977).
Reproductive phenology varies between species with different life-forms (see Appendix 1).
Species in the different genera of globose cacti, such as Mammillaria, Sclerocactus, and
Turbinicarpus (Martinez et al., 1994; Lockwood, 1995; Contreras & Valverde, 2002; Valverde
et al., in press), begin to reproduce in early winter, and their reproductive periods last for more
than 5 months. In contrast, species in the different genera of columnar cacti may reproduce in
autumn/winter (Cereus, Myrtillocactus, Pachycereus, Pilosocereus, Selenicereus, Stenocereus) or
spring/summer (Carnegiea, Cephalocereus, Escontria, Hylocereus, Lophocereus, Neobuxbaumia,
Pachycereus, Pilosocereus, Polaskia, Stenocereus, Subpilocereus), and their reproductive periods
last for 2 to 4 months (Steenbergh & Lowe, 1977; Weiss et al., 1994; Valiente-Banuet et al., 1996,
1997a, 1997b; Nassar et al., 1997; Casas et al., 1999; Rojas-Martinez et al., 1999; Ruiz et al., 2000;
Fleming et al., 2001; Esparza-Olguin et al., 2002). Other columnar cacti, however, may produce
flowers and fruitsthroughoutthe year (Praecereus, Stenocereus, Weberbaurocereus, Weberocereus;
Sahley, 1996; Nassar et al., 1997; Tschapka et al., 1999; Ruiz et al., 2000). Species in the genus
Opuntia produce flowers and fruits in spring/summer (0. brunneogemmia, 0. compressa, 0. dis-
cata, 0. imbricata,0. lindheimeri,0. phaecantha,0. polyacantha,0. rastrera,0. stricta,0. viridirubra; Grant et al., 1979; Spears, 1987; Osbom et al., 1988; McFarland et al., 1989;
Mandujano et al., 1996; Schlindwein & Wittmann, 1997) or in winter (0. spinosissima; Negr6nOrtiz, 1998). Finally, barrel cacti in the genera Echinocereus, Echinomastus, and Ferocactus
reproduce mainly during spring/summer (Breckenridge & Miller, 1982; Johnson, 1992; McIntosh, 2002).
Field observations regarding flower and fruit production in some cactus species (Carnegiea
gigantea, Echinomastuserectrocentrus,Lophocereusschottii,Mammillariacrucigera,M mag-
186
I3-D
'50
.3
ID
2C
1
20
O~~~~~~~~~
-46
6-
r~~~~~~~~~~~~~~""
6 4
1 2 34567E
Size Category
Fig. 1. Examples of the relationship between fecundity and size for cacti with different life-forms.
a. The globose cactus Mammillariacrucigera. b. The columnar cactus Neobuxbaumiatetetzo. c. The
platyopuntia Opuntia rastrera.
IN THE CACTACEAE
DEMOGRAPHY
187
nimamma,Neobuxbaumiamacrocephala,N. tetetzo,Pachycereuspringlei,Stenocereusqueretaroensis, S. thurberi;Steenbergh& Lowe, 1977; Johnson, 1992; Johnsonet al., 1992; Pimienta
et al., 1998; Godinez-Alvarezet al., 1999; Fleming et al., 2001; Contreras& Valverde,2002;
Esparza-Olguinet al., 2002; Valverde et al., in press) have shown that there is interannual
variationin the timing and intensityof reproductiveevents. These variationsmay be relatedto
particularenvironmentalfactors, such as the presence of winter rainfall,the early startof the
rainyperiod,and/ora high total annualprecipitation(Bowers, 1996b;Valverdeet al., in press).
The numberof fruitsproducedper plant and the numberof seeds per fruitvary according
to species and life-form (Table III). For instance, fruits of columnarcacti may producemore
than 1000 seeds, whereas fruitsof globose cacti generallyproducefewer than 100 seeds. The
numberof seeds per fruit in barrelcacti, cylindropuntias,and platyopuntiasvaries between
10 and 200.
In additionto sexual reproduction,many cacti are capableof spreadingvegetatively(Parker
& Hamrick, 1992; Mandujanoet al., 1998). This ability is common in the genus Opuntia,
although plants in other genera, such as Lophocereus, Myrtillocactus, Pachycereus, and
Stenocereus,may also spreadthroughrametproduction(pers. obs.). The relativeimportanceof
vegetative spreadcomparedwith sexual reproductionis relatedto differentfactorssuch as soil
type, the presence of nurse plants, herbivory,and flooding (Mandujanoet al., 1998). Opuntia
engelmanniplants apparentlyadjustthe relative proportionof new cladodes and flowers in
responseto the previousyear's resourceexpenditure(Bowers, 1996c).
VI. Size Structure
Population structure is the result of size- (or age-) specific birth and mortality rates
(Silvertown, 1987). The particularsize- (or age-) structurefound in a populationmay be determined by several extrinsic and intrinsicfactors.Among these, seed production,germination
rates, soil seed banks, vegetative spread,herbivory,and weatherpatternsmay play an important role (Silvertown, 1987). Histogramsrepresentingpopulationstructurefor differentcactus
species often show an uneven distributionof individualsamong size classes (Fig. 2; Appendix
1), which suggests that recruitmentoccurs in pulses, apparentlyassociated with infrequent
favorable conditions for germinationand establishment(Mandujanoet al., 2001; EsparzaOlguin et al., 2002). These favorableconditionsappearto be relatedto particularprecipitation
andtemperaturecombinationsfor species suchas Carnegieagigantea,Echinocactushorizonthalonius, Ferocactus cylindraceus,Neobuxbaumiamacrocephala,and Opuntiaechios (Reid et
al., 1983; Bowers, 1997b;Pierson& Turner,1998; Hicks & Mauchamp,2000; Esparza-Olguin
et al., 2002). As noted above, recruitmentpulses of some species may occur in rainy years
associatedwith particularclimatic events such as El Ninfto(Hicks & Mauchamp,2000). In the
case of Opuntiarastrera,germinationand establishmentare enhancedwhen nurse plants are
readily availableand herbivoryis low (Mandujanoet al., 1998).
Othercactusspecies show a type of populationstructurein which the numberof individuals
decreasesmonotonicallywith size or age (i.e., Ferocactusacanthodes,Neobuxbaumiatetetzo,
Opuntiaechios; Jordan& Nobel, 1981; Godinez-Alvarezet al., 1999; Hicks & Mauchamp,
2000; Fig. 2; Appendix 1). This type of populationstructuresuggests growingpopulationsthat
areregeneratingconstantly,in which mortalityratedecreaseswith size or age (Martinez-Ramos
& Alvarez-Buylla,1995).
Populationsthat spread throughouta variety of environmentalconditions, such as those
occupying altitudinalgradients,may show distinct populationstructuresin differenthabitat
types. This is the case in Opuntiaechios, in which populationsin a low arid habitatshow a
188
Table III
Mean number of seeds per fruit and standard errors for species of cacti with different life-forms
Species
Columnarcacti
Carnegieagigantea
Cereusperuvianus
Lophocereusschottii
Neobuxbaumiamacrocephala
N. mezcalaensis
N. tetetzo
Pachycereuspringlei
Pilosocereus moritzianus
Stenocereusgriseus
S. gummosus
S. stellatus
S. thurberi
Subpilocereushorrispinus
S. repandus
Weberbaurocereusweberbaueri
Barrel cacti
Echinomastuserectrocentrus
Melocactus violaceus
Globose cacti
Mammillariacrucigera
M magnimamma
Cylindropuntias
Opuntia imbricata
Platyopuntias
Opuntia brunneogemmia
0. engelmannii
Opuntiaphaeacantha
0. polyacantha
O. rastrera
0. stricta
0. viridirubra
Seeds
per fruit
Standard
error
2263
1358
1241.8
1349.8
1946.7
1952
126
465
552
496
933
1288.1
1330.4
1329.1
2496
1121
1581
674
750
934
1969
536.9
351.3
1123
1483
1067
108
140
112.7
99.3
131.8
6.3
209
30
32.9
51.1
91.7
162.2
141.1
165.3
91
62.2
99.8
15
35
161.3
101.5
77.3
51.6
93
-
86.6
21.8
20
93.48
93.5
Number
of fruits
sampled
Source
18
15
25
51
44
10
168
25
10
10
35
23
15
18
15-27
27
15-27
25
20
20
19
28
8
15-27
15-27
14
Nassar et al., 1997

Nassar et al., 1997
Sahley, 1996
4.4
1.6
16
Johnson, 1992
Figueira et al., 1994
12.6
36.8
42
112
69
69
Contreras& Valverde,2002
Valverdeet al., in press
62
8.4
36
McFarlandet al., 1989
55.2
143.9
171.6
5.8
21.5
16.8
20
20
20
Schlindwein & Wittmann,1997

Bowers, 1997a
30
30
31
30
36
-
Osbom et al., 1988
50.2
72.7
25.1
9.0
208
60.6
65.3
67.5
63.7
6.9
5.7
3.6
2.5
0.13
3.7
4.8
4.5
3.7
Steenbergh& Lowe, 1977

Fleming et al., 2001
Silva, 1988
Parker,1989
Esparza-Olguinet al., 2002
Valiente-Banuetetal., 1997a
Valiente-Banuetet al., 1997a
Godinez-Alvarez et al., 2002
Nassar et al., 1997

Silvius, 1995
Nassaret al., 1997
Le6n de la Luz & Cadena, 1991
Casas et al., 1999
Parker,1987
Osbom et al., 1988

Mandujanoet al., 1996
Spears, 1987
20
Schlindwein & Wittmann,1997
DEMOGRAPHY IN THE CACTACEAE

026
0.3 -
ab
0e.16 1
0.26
0.2
189
0 16X10-
5.'
S6S
910
626 .0.36
56
02
c~~~~~~~~~~~2
0.16
02-f
01 6-
.3
'
B67
1
Size
3'
Cago.y
Fig. 2. Populationsize structures.a. The barrelcactus Echinocactushorizonthalonius.b. The globose

cactus Mammillariacrucigera. c. The columnarcactus Neobuxbaumiamacrocephala.d. The columnar
cactus Neobuxbaumiatetetzo.
smooth decline in the numberof individualsfrom small to large size classes, whereaspopulations at higher altitudesshow a bimodal structurewith few plants of intermediatesizes (Hicks
& Mauchamp,2000).
VII. Population Dynamics

With the increasingpopularityof projectionmatrix models, plant populationecology has
grown significantlyin recent decades. However, most of the populationsfor which projection
matriceshave been constructedcorrespondto herbaceousspecies fromtemperatehabitats(e.g.,
Bierzychudek,1982; Valverde& Silvertown, 1998). Thereare only a few referencesfor cactus
populationsthat include the use of matrixmodels. Yet the informationavailableis quite valuable, for it includes cactus species with differentgrowth forms for which demographicinformationhas been gatheredduringseveralyears and/orat various locations (TableIV). For each
species, we considerthe matricesof differentyears and/orsites separately,insteadof constructing one average matrix per species. We believe that this approachallows us to analyze the
variationin demographicbehaviorboth within species and between species. If averagematrices were built, the within-speciesvariationwould be totally obscured,and flawed conclusions
could arise regardingthe differencesbetween species.
Fromthe informationpresentedin TableIV, it may be noted thatpopulationgrowthrate(A)
is close to unity for most records;only in two populationsis it significantly lower than unity
(Carnegiea gigantea and Mammillaria crucigera 2), and only in one population is it signifi-
cantly largerthanunity (Mammillariamagnimamma2). This suggest thatmost cactus populations studiedare close to the numericalequilibrium.Yet annualprojectionmatricesmay be a
limitedtool for detecting long-termincreasesor declines in populationnumbers,especially in
Table IV
Summary of population dynamics data for 17 cactus populations of eight specie

Population growth rates (A) were calculated from projection matrices from n individuals each. Elasticity valu
entries representing the seed bank (el), sexual reproduction via seed and/or seedling production (e2), clonal
stasis (e6) and growth (e6). Elasticity values per demographic process are reproduction (R), surviva
and per life-cycle stage are seedlings (S), juveniles (J), and adults (A).
Elasticity values
Per demographic proces

Species
Carnegia gigantea
Neobuxbaumiatetetzo
Neobuxbaumiamacrocephala a
1
2
Escontria chiotilla b
1
2
Mammillariamagnimammac
1
2
3
4
Mammillariacrucigeraa
1
2
Coryphantarobbinsorumd
1
2
3
Opuntiarastrera d
1
2
el
e2
e3
e4
0.540
15
1.000 1900
0
0.009
0.979
0.994
206
206
0.006
0.006
1.011
1.04
400
400
0.020
0.025
0.956
1.333
0.967
0.945
203
203
206
206
0.003
0.207
2E-05
0.006
0.015
0.019
7E-06
0.125
0.977
0.896
230
230
0.001
0.002
0.002
0.006
1.093
1.006
1.042
63
39
63
0.059
0.018
0.058
1.04 3000
0.997 250
0.001
1E-04
0.014
0.001
0.052
0.021
0
0.05
0.100
0.050
e5
e6
0.99
0.914
0
0.078
0
0.000
0.99
0.914
0
0.078
0.837
0.885
0.105
0.088
0.006
0.006
0.889
0.906
0.105
0.088
0.791
0.763
0.189
0.212
0.020
0.025
0.791
0.763
0.189
0.212
0.934 0.044
0.334 0.436
0.999 7E-04
0.713 0.156
0.003
0.207
2E-05
0.006
0.948
0.353
0.999
0.838
0.044
0.436
0.001
0.156
0.984
0.971
0.013
0.021
0.001
0.002
0.986
0.977
0.013
0.021
0.762
0.927
0.769
0.179
0.054
0.173
0.059
0.018
0.058
0.762
0.927
0.769
0.178
0.054
0.173
0.570
0.780
0.280
0.150
0.001
0.020
0.670
0.830
0.330
0.150
Sources: 1 = Steenbergh& Lowe, 1977, in Silvertown et al., 1993; 2 = Godinez-Alvarez et al., 1999; 3 = Esparza-Olguine
5 = Valverdeet al., in press; 6 = Contreras& Valverde, 2002; 7 = Schmalzel et al., 1995; 8 = Mandujanoet al., 2001.
a One population in each case, sampled in different years. b Two different sites sampled within the same time
period. c Two
dEach record correspondto a different site, and in each case the average results of several years is given.
IN THE CACTACEAE
DEMOGRAPHY
191
long-lived species in which relevantpopulationprocesses may take place at the scale of decades (Pierson& Turner,1998; de Kroon et al., 2000).
As noted in previous sections, the analysis of transitionmatricesrevealed that mortality
decreases and reproductionincreases with increasingplant size in all species. Additionally,
plant growthrates are slow, which may be noted by consistentlylow transitionand high "stasis"probabilitiesin most populationmatrices.Only the matricesfor Mammillariaspecies show
rapidgrowthtransitions,in which plantsmay grow more thanone size categoryfrom one year
to the next. In some species "regressions"to smallersize categoriesare possible due to branch
death,particularlyin globose species. Among the species analyzed,vegetative spreadoccurs
only in Opuntiarastrera, although,as noted above, other species may also show this ability.
Anotheraspect that may be noticed from the analyzedprojectionmatricesis the almost total
absence of a seed bank in the models, with the exception of 0. rastrera.Although it appears
that the seeds of various cactus species have the ability to remainviable in the soil for more
than a year (see section IV), it is believed that most seeds either germinate(given their high
germinability)or die shortlyafterthey are shed.
A. ELASTICITY
ANALYSES
The relative contributionof differenttransitionsto the value of X has traditionallybeen
evaluated through elasticity analyses (de Kroon et al., 1986). In additionto evaluatingthe
elasticityof individualmatrixentries,it is possible to addup elasticityvalues of severalmatrix
entriescorrespondingto the same demographicprocess in orderto analyze the relativecontributionof each process to populationgrowthrate(Silvertownet al., 1993). Herewe subdivided
the life cycle of cactusspecies into six differenttypes of transitions,following Silvertownet al.
(1993): persistenceof seeds in the seed bank (el), sexual reproduction(e2), clonal propagation
(e3), retrogressionto smaller size classes (e4), persistenceor "stasis"in the currentsize class
(e5),and growthto largersize classes (e6).TableIV shows the values of e,-e6 for the 17 matrices (correspondingto eight species) availablein the literaturefor cactus species. Severaltrends
are apparentfromthese data:as noted above, seed banksandvegetativepropagationare absent
in most cases (with the exceptionof Opuntiarastrera2); sexual reproductioncontributesvery
little to populationgrowth rate; and the most importantcontributionsare in e5, which correspondto the persistenceof individualsin theircurrentsize class. These trendsare also apparent
when the differentelasticityvalues were addedup to accountfor the three main demographic
processes consideredby Silvertownet al. (1993): fecundity (e} + e2), survival (e4 + e5), and
growth (e3 + e6) (Table IV). Thus, plotting the differentpopulationsin the demographictriangle, we found that most of them fell towardthe survivalend (Fig. 3, top), which coincides
with the patternobservedfor long-lived perennials(Silvertownet al., 1993). Only one population, Mammillariamagnimamma2, fell closer to the center of the triangle,which implies a
balance between the contributionof growthand survivaland a relativelyhigh contributionof
fecundityto populationgrowthrate.This populationis the one thatexhibitsthe largestX. Thus,
as observed by other authors(Valverde& Silvertown, 1998; de Kroon et al., 2000), those
populationswith largerXs tend to be locatedcloser to the centerof the triangle,whereaslower
As generallyoccupy the far-rightcornerof the triangle(Fig. 4, top).
The position of the differentpopulationsin the demographictriangledoes not seem to be
related to either life-form or longevity, as was suggested by Rosas-Barreraand Mandujano
(2002). For instance, the Carnegieagigantea and the Mammillariamagnimamma3 populations (a columnarlong-lived species and a globose shorter-livedspecies, respectively)are locatedat the far-rightcornerof the triangle,almostin the vortex,while those populationslocated
closer to the centerof the triangleincludeEscontria chiotilla 2, Coryphantharobbinsorum1,
Opuntiarastrera 1, and Mammillariamagnimamma2, representingthree differentlife-forms.
192
Cl41v,o
ltoll
'
''
oo,~~~~~~~~~~~~~0
,.|/'
1/
f
-1,0
"44. 1
>
__
,y
.-04
0.0
)
*t.ll~~~~~~~~~~o
C[
02
0,0
_~~~~~~~~~~~~~~~)
'
seedling
Fig. 3 (above). Triangularordinationof elasticityvalues for the 17 matricesreportedin TableIV. The

three axes correspondto fecundity,growth, and survival (top) and seedlings, juveniles, and adults (bottom). See the text for furtherexplanation.
Fig. 4 (facing page). Elasticityvalues for the 17 matricesreportedin TableIV ordainedon a triangular plot representingdifferentdemographicprocesses (top) and differentlife-cycle stages (bottom),along
with the A value for each matrixrepresentedin the y axis.
193
i2
I,....
BA-. ,.~i
1.04
:....ft.'
AC'
0~~~~~~~~~~~~~~7
N.
'4.
194
In additionto evaluatingthe contributionof particulardemographicprocesses, as proposed

by Silvertownet al. (1993), it is possible to use elasticityvalues to analyzethe contributionof
differentlife-cycle phases to populationgrowthrate.Here we have consideredthreelife-cycle
phases: seedlings (newly establishedindividuals,generallycorrespondingto the first and, occasionally,also the second and the thirdsize classes);juveniles (categoriesof well-established
individuals,with mortalityprobabilitieslower thanthose of seedlings but no sexual reproduction); and adults(reproductiveindividualsbelongingto largersize classes) (TableIV). In most
of the cactus populationswe detected a relatively largercontributionof adults to population
growth rate, comparedwith those of seedlings and juveniles (Fig. 3, bottom). Only the two
Escontria chiotilla populations(located towardthe top of the trianglein Fig. 3, bottom)show
a high contributionof juveniles to X, whereasMammillariamagnimamma2 and Coryphantha
robbinsorum1 show also a substantialcontributionfrom seedlingsto X.The latterhigh elasticity values for the seedlingphasecoincide with the largestks (Fig. 4, bottom).This patternis not
particularlyclear,however,since some populationswith very low X values have relativelyhigh
seedling-elasticityvalues (i.e., Carnegiea gigantea) and some populationswith a relatively
high Kshow very low elasticityvalues for the seedling phase (TableIV; Fig. 4, bottom).
B. MATRIX
MODELSANDMATRIXSIMULATIONS
In some of the demographicstudiescompiled in TableIV, authorsalso reportthe resultsof
matrix simulationexercises in which particularmatrix entries were modified to explore the
potentialeffect of these changeson populationgrowthrate.A commonresultof these exercises
is that a substantialincreasein fecunditydoes result in an increasein populationgrowthrate,
despitethe fact thatfecundityentriesgenerallyshow low elasticities,whereaschangesin other
matrixentries generallyhave a low impacton K (Schmalzel et al., 1995; Ortega-Baes,2001;
Contreras& Valverde,2002; Esparza-Olguinet al., 2002).
Mandujanoet al. (2001) combinedthe demographicinformationfor Opuntiarastrerafor
eight years and at two sites in orderto generatedifferentprojectionmodels; that is, the time
invariant(most generally used) yearly matrices,average matricesfor several years, periodic
matrices, and stochastic projections.They conclude that the X values obtained using these
differentmethods are quite similar.However,they assertthat averagematricesare not an adequate way of exploring environmentalheterogeneity,because they do not take into account
variabilityin demographicbehavioracrossyears.In this case the authorsrecommendthe use of
periodicmatricesbecausethey integrateinterannualvariationandgive more consistentelasticity values (Mandujanoet al., 2001). Stochasticsimulationshave also been used in Mammillaria magnimamma(Valverdeet al., in press). The authorssuggest that this type of approach
may be useful in projectinglong-termpopulationdynamicsof cactus species, especially as an
aid to understandingnot only long-termpopulationgrowthrates but also transientdynamics.
This approachmay be especially fruitfulwhen the probabilityof occurrenceof environmental
phenomenathat most dramaticallyaffect mortalityand recruitmentare known and can be incorporatedin the stochasticprojection(Valverdeet al., in press).
VIII. Conclusions and Perspectives
The review presentedhere leaves us with the certaintythat cactus populationecology is
definitely a consolidatingsubject within plant ecology. The numberof publicationsand the
amountof ecological informationon the subjectis alreadysubstantialand is growing day by
day.Yet certaintopics are clearlymore developedthanothers.The ecology of seed germination
is perhapsone of the subjectsto which greaterattentionhas been directed,althoughwe still
DEMOGRAPHY
IN THE CACTACEAE
195
lack informationaboutthe potentialrole of seed banks. In recentyears the nurse-cactusassociation has also been the subject of a substantialamountof work; however, we need further
researchin orderto evaluatethe importanceof this phenomenonfor cacti with differentlifeforms. On the other hand, analysis of cactus populationdynamics throughmatrixmodels is
ratherincipient. On this same topic, the relative importanceof sexual reproductionversus
vegetativepropagationin the maintenanceof those populationsthatmay show clonal growthis
still unknown.
In regardto the differentlife-forms representedin the Cactaceae,columnarcacti have received the greatestattention;ecological researchon the globose and the barrelcacti, as well as
the cylindropuntiasand platyopuntias,has been less intense.In addition,most of the literature
refersto NorthAmericanandCentralAmericancacti, and little is knownaboutSouthAmerican
species.
In exploringthe literatureon cactus populationecology, it has been interestingto discover
that biotic interactionssuch as nurse-cactus associations, pollination and seed dispersal by
animals,seed predation,herbivory,and competition,appearto be importantorganizingforces
in desert and semi-desertcommunities(Valiente-Banuet& Godinez-Alvarez,2002; ValienteBanuetet al., 2002). Previouswork on desertecology had failed to recognizethis aspect, since
it was believed thatthe main organizingforces of communitiesin stressfulhabitatswere limiting abiotic factors (Noy-Meir, 1973, 1974, 1979/1980; Grime, 1979). However,the evidence
compiledin this review suggests thatbiotic forces may exertdramaticeffects on desertpopulations, thereforefunctioningas key factorsdeterminingcommunitycompositionand structure.
We conclude with a reflection on cactus conservation.As we mentionedbefore, it is well
known thatmany cactus species are vulnerableor in dangerof extinction.Yet, of all the cactus
species for which any kind of populationecology informationis available, only a few have
been classified as rare, vulnerable,or endangered(see Appendix 1). Only through a better
understandingof populationprocesses will we be able to account for ecological phenomena
such as rarityand populationdecline, which are widespreadamong endangeredplants. Thus
we must intensifyour effortsto deepen ourunderstandingof cactuspopulationdynamicsif we
are to offer any tools with which to preservethe biodiversityof this fascinatingplant family.
IX. Acknowledgments
We are grateful to M. C. Mandujanofor supplying us with a copy of Rosas-Barrera&
Mandujano(2002) even before it was published.We thankM. Francoand J. Silvertownfor
allowing us to use the matrixthey compiled for Carnegieagigantea (used in Silvertownet al.,
1993) from previouslypublishedwork. We are also gratefulto A. Valiente-Banuet,who supervised the work of P. Ortega-Baeson Escontria chiotilla.
X. Literature Cited
Altesor, A., E. Ezcurra & C. Silva. 1992. Changes in the photosyntheticmetabolismduringthe early
ontogeny of four cactus species. Acta Oecol. 13: 777-785.
Anderson, E. F., S. Arias Montes & N. P. Taylor. 1994. Threatenedcacti of Mexico. Royal Botanical
Gardens,Kew, England.
Arriaga, L., Y. Maya, S. Diaz & J. Cancino. 1993. Association between cacti and nurseperennialsin a
heterogeneoustropicaldry forest in northwesternMexico. J. Veg. Sci. 4: 349-356.
Bierzychudek, P. 1982. Life historiesand demographyof shade-toleranttemperateforestherbs:A review.
New Phytol. 90: 757-776.
Bowers, J. E. 1996a. Growth rate and life span of a prickly pear cactus, Opuntiaengelmannii, in the
northernSonorandesert. Southw.Naturalist41: 315-318.
196
THE BOTANICALREVIEW
. 1996b. Environmentaldeterminantsof floweringdate in the columnarcactus Carnegieagigantea

in the SonoranDesert. Madrofio43: 69-84.
. 1996c. More flowers or new cladodes?Environmentalcorrelatesand biological consequencesof
sexual reproductionin a SonoranDesertpricklypearcactus, Opuntiaengelmannii.Bull. TorreyBot.
Club 123: 34-40.
. 1997a. The effect of droughton Engelmannpricklypear (Cactaceae:Opuntiaengelmannii)fruit
and seed production.Southw.Naturalist42: 240-242.
. 1997b. Demographicpatternsof Ferocactuscylindraceusin relationto substrateage and grazing
history.P1.Ecol. 133: 37-48.
. 2000. Does Ferocactus wislizeni (Cactaceae)have a between-yearseed bank?J. Arid Environm.
45: 197-205.
, R. H. Webb & R. J. Rondeau. 1995. Longevity,recruitmentand mortalityof desert plants in
GrandCanyon,Arizona,USA. J. Veg. Sci. 6: 551-564.
Bravo-Hollis, H. & H. Sanchez-Mejorada. 1978. Las Cactaceasde Mexico. Vol. 1. UniversidadNacional
Aut6nomade Mexico, Mexico City.
. 1991. Las Cactaceasde Mexico. Vols.2-3. UniversidadNacionalAut6nomade Mexico,
&
Mexico City.
Breckenridge, F. G, III & J. M. Miller. 1982. Pollinationbiology, distribution,and chemeotaxonomyof
the Echinocereusenneacanthuscomplex (Cactaceae).Syst. Bot. 7: 365-378.
Briones, O., C. Montania& E. Ezcurra. 1998. Competitionintensityas a functionof resourceavailability in a semiaridecosystem. Oecologia 116: 365-372.
Callaway, R. M. 1995. Positive interactionsamong plants. Bot. Rev. (Lancaster)61: 306-349.
. 1998. Are positive interactionsspecies-specific?Oikos 82: 202-207.
Carrillo-Garcia, A., Y. Bashan & G J. Bethlenfalvay. 2000. Resource-islandsoils and the survivalof
the giant cactus, cardon,of Baja CaliforniaSur. P1.& Soil 218: 207-214.
Casas, A., A. Valiente-Banuet, A. Rojas-Martinez & P. Davila. 1999. Reproductivebiology and the
process of domesticationof the columnarcactus Stenocereusstellatus in centralMexico. Amer. J.
Bot. 86: 534-542.
Cody, M. L. 1993. Do cholla (Opuntiaspp., Subgenus Cylindropuntia)use or need nurse plants in the
Mojave Desert? J. Arid Environm.24: 139-154.
. 2000. Slow-motionpopulationdynamicsin MojaveDesertperennialplants.J. Veg. Sci. 11:351-358.
Contreras, C. & T. Valverde. 2002. Evaluationof the conservationstatusof a rarecactus (Mammillaria
crucigera) throughthe analysis of its populationdynamics.J. Arid Environm.51: 89-102.
De Kroon, H., H. Plaiser, J. van Groenendael & H. Caswell. 1986. Elasticity:The relativecontribution
of demographicparametersto populationgrowth rate. Ecology 67: 1427-1431.
I ~~ & J. Ehrlen. 2000. Elasticities:A review of methodsand model limitations.Ecology 81:
607-618.
De Viana, M. L. 1996-1997. Distribuci6nespacial de Trichocereuspasacana (Cactaceae)en relaci6n al
espacio disponibley al banco de semillas. Revista Biol. Trop.44/45: 95-103.
, N. Acreche, R. Acosta & L. Morafna.1990. Poblaci6ny asociacionesde Trichocereuspasacana
(Cactaceae)en Los Cardones,Argentina.Revista Biol. Trop. 38: 383-386.
, S. Suhring & B. F. J. Manly. 2001. Applicationof randomizationmethodsto studythe association of Trichocereuspasacana (Cactaceae)with potentialnurse plants. PI. Ecol. 156: 193-197.
Dubrovsky, J. G 1996. Seed hydrationmemory in SonoranDesert cacti and its ecological implication.
Amer.J. Bot. 83: 624-632.
. 1998. Discontinuoushydrationas a facultativerequirementfor seed germinationin two cactus
species of the SonoranDesert. J. TorreyBot. Soc. 125: 33-39.
Esparza-Olguin, L., T. Valverde & E. Vilchis-Anaya. 2002. Demographicanalysis of a rarecolumnar
cactus (Neobuxbaumiamacrocephala)in the TehuacanValley, Mexico. Biol. Con. 103: 349-359.
Figueira, J. E. C., J. Vasconcellos-Neto, M. A. Garcia & A. L. Teixeira de Souza. 1994. Saurocoryin
Melocactus violaceus (Cactaceae).Biotropica 26: 295-301.
Fleming, T. H. & A. Valiente-Banuet (eds.). 2002. Columnarcacti and their mutualists:Evolution,
ecology, and conservation.Universityof ArizonaPress, Tucson.
, C. T. Sahley, J. N. Holland, J. D. Nason & J. L. Hamrick. 2001. SonoranDesert columnar
cacti and the evolution of generalizedpollinationsystems. Ecol. Monogr.71: 511-530.
197
Flores, F. J. L. & R. I. Yeaton. 2000. La importanciade la competencia en la organizaci6n de las

comunidadesvegetales en el altiplanomexicano. Interciencia(Caracas)25: 365-371.
Flores, J. & 0. Briones. 2001. Plantlife-formand germinationin a Mexican inter-tropicaldesert:Effects
of soli water potentialand temperature.J. Arid Environm.47: 485-497.
Flores-Martinez, A., E. Ezcurra & S. Sanchez-Col6n. 1994. Effect of Neobuxbaumiatetetzoon growth
and fecundityof its nurse plant Mimosa luisana. J. Ecol. 82: 325-330.
&
. 1998. Wateravailabilityand the competitiveeffect of a columnarcactuson its
nurseplant.Acta Oecol. 19: 1-8.
Franco, A. C. & P. S. Nobel. 1989. Effect of nurseplantson the microhabitatand growthof cacti. J. Ecol.
77: 870-886.
Garcia-Moya, E. & M. C. McKell. 1970. Contributionof shrubsin the economy of a desert-washplant
community.Ecology 51: 81-88.
Gibson, A. C. & P. S. Nobel. 1986. The cactus primer.HarvardUniv. Press. Cambridge,MA.
Godinez-Alvarez, H. & A. Valiente-Banuet. 1998. Germinationand early seedling growthof Tehuacan
Valley cacti species: The role of soils and seed ingestion by disperserson seedling growth. J. Arid
Environm.39: 21-31.
~~ & B. L. Valiente. 1999. Biotic interactionsand the populationdynamics of the longlived columnarcactus Neobuxbaumiatetetzo in the TehuacanValley, Mexico. Canad. J. Bot. 77:
203-208.
& A. Rojas-Martinez. 2002. The role of seed dispersersin the populationdynamicsof
the columnarcactus Neobuxbaumiatetetzo. Ecology 83: 2617-2629.
Goldberg, D. E. & R. M. Turner. 1986. Vegetationchange and plant demographyin permanentplots in
the SonoranDesert. Ecology 67: 695-712.
Grant, V., K. A. Grant & P. D. Hurd Jr. 1979. Pollinationof Opuntialindheimeriand relatedspecies.
P1. Syst. Evol. 132: 313-320.
Grime, J. P. 1979. Plant strategiesand vegetationprocesses. Wiley, Chichester.
Gulmon, S. L., P. W. Rundel, J. R. Ehleringer & H. A. Mooney. 1979. Spatialrelationshipsand competition in a Chileandesert cactus. Oecologia 44: 40-43.
Haman, 0. 2001. Demographicstudies of three indigenousstand-formingplant taxa (Scalesia, Opuntia,
and Bursera) in the GalapagosIslands,Ecuador.Biodiv. & Conserv. 10: 223-250.
Hastings, J. R. & S. M. Alcorn. 1961. Physical determinationsof growthand age in the giant cactus. J.
Ariz. Acad. Sci. 2: 32-39.
Hernandez, H. M. & H. Godinez-Alvarez. 1994. Contribuci6n al conocimiento de las Cactaceas
mexicanasamenazadas.Acta Bot. Mex. 26: 33-52.
Hicks, D. J. & A. Mauchamp. 2000. Populationstructureand growth patternsof Opuntiaechios var.
gigantea along an elevationalgradientin the GalapagosIslands. Biotropica32: 235-243.
Horvitz, C. C. & D. W. Schemske. 1995. Spatiotemporalvariationin demographictransitionsof a tropical understoreyherb: Projectionmatrixanalysis. Ecol. Monogr.65: 155-192.
Huerta, F. M. & V. E. Escobar. 1998. Estatusecol6gico actual de Ferocactus hystrix(DC) Lindsay en
los Llanos de Ojuelos, Jalisco-Zacatecas.Cact. Suc. Mex. 43: 57-64.
Hunt, D. (comp.). 1999. C.I.T.E.S.Cactaceaechecklist. Ed. 2. Royal Botanic Gardens& International
Organizationfor SucculentPlant Study,Kew, England.
Hutto, R. L., J. R. McAuliffe & L. Hogan. 1986. Distributionalassociates of the saguaro(Carnegiea
gigantea). Southw.Naturalist31: 469-476.
Johnson, R. A. 1992. Pollinationand reproductiveecology of Acufiacactus,Echinomastuserectrocentrus
var. acunensis (Cactaceae).Int. J. P1. Sci. 153: 400-408.
, M. A. Baker, D. J. Pinkava & G A. Ruffner. 1992. Seedling establishment,mortality and
flower productionof the Acufia cactus, Echinomastuseretrocentrusvar. acunensis (Cactaceae).Pp.
170-180 in R. Sivinski & K. Lightfoot (eds.), Southwesternrare and endangeredplants: Proceedings of the SouthwesternRare and EndangeredPlant Conference.New Mexico Forestryand Resources ConservationDivision, Energy,MineralsandNaturalResourcesDepartment,SantaFe, NM.
Jordan, P. W. & P. S. Nobel. 1981. Seedling establishmentof Ferocactus acanthodes in relation to
drought.Ecology 62: 901-906.
&
. 1982. Height distributionof two species of cacti in relationto rainfall,seedling establishment,and growth. Bot. Gaz. 143: 511-517.
9
198
THE BOTANICALREVIEW
Larmuth,J. & H. J. Harvey. 1978.Aspectsof the occurrenceof desertplants.J.AridEnvironm.1: 129-133.

Leirana-Alcocer, J. & V. Parra-Tabla. 1999. Factorsaffectingthe distribution,abundanceand seedling
survivalof Mammillariagaumeri, an endemic cactus of coastalYucatan,Mexico. J. Arid Environm.
41: 421-428.
Le6n de la Luz, J. L. & R. D. Cadena. 1991. Evaluaci6nde la reproducci6npor semilla de la pitaya
agria (Stenocereusgummosus)en Baja CaliforniaSur, Mexico. Acta Bot. Mex. 14: 75-87.
Lockwood, M. W. 1995. Notes on life historyof Ancistrocactustobuschii(Cactaceae)in Kinney County,
Texas. Southw.Naturalist40: 428-430.
Mandujano, M. C., C. Montafia & L. Eguiarte. 1996. Reproductiveecology and inbreedingdepression
in Opuntiarastrera(Cactaceae)in the ChihuahuanDesert: Why are sexually derivedrecruitments
so rare?.Amer.J. Bot. 83: 63-70.
,I. Mendez & J. Golubov. 1998. The relative contributionsof sexual reproductionand
clonal propagationin Opuntiarastrera from two habitats in the ChihuahuanDesert. J. Ecol. 86:
911-921.
,IM. Franco, J. Golubov & A. Flores-Martinez. 2001. Integrationof demographic
annualvariabilityin a clonal desert cactus. Ecology 82: 344-359.
Martinez, D., A. Flores-Martinez, F. L6pez & G Manzanero. 2001. Aspectos ecol6gicos de Mammillaria oteroi Glass & R. Foster en la regi6n mixteca de Oaxaca,Mexico. Cact. Suc. Mex. 46: 32-39.
Martinez, J. G, H. Suzian & C. A. Salazar. 1993. Aspectos ecologicos y demograficosde Ariocarpus
trigonus(Weber)Schumann.Cact. Suc. Mex. 38: 30-38.
&
. 1994. Aspectos ecol6gicos y demograficosde Neolloydia pseudopectinata
(Backeberg)E. F. Anderson.Cact. Suc. Mex. 39: 27-33.
Martinez-Ramos, M. & E. Alvarez-Buylla. 1995. Ecologia de poblaciones de plantas de una selva
humedade Mexico. Bol. Soc. Bot. Mexico 56: 121-153.
McAuliffe, J. R. 1984a. Sahuaro-nursetree associations in the SonoranDesert: Competitiveeffects of
sahuaros.Oecologia 64: 319-321.
. 1984b. Prey refugia and the distributionsof two sonorandesert cacti. Oecologia 65: 82-85.
& F. J. Janzen. 1986.Effectsof intraspecificcrowdingon wateruptake,waterstorage,apicalgrowth,
and reproductivepotentialin the sahuarocactus, Carnegieagigantea. Bot. Gaz. 147: 334-341.
McFarland, J. D., P. G Kevan & M. A. Lane. 1989. Pollinationbiology of Opuntiaimbricata(Cactaceae)
in southernColorado.Canad.J. Bot. 67: 24-28.
McIntosh, M. E. 2002. Floweringphenology and reproductiveoutputin two sister species of Ferocactus
(Cactaceae).PI. Ecol. 159: 1-13.
Meyran, J. 1980. GuiabotAnicade Cactaceasy otrassuculentasdel Vallede Tehuacan.SociedadMexicana
de CactologiaA.C., Mexico City.
Mourelle, C. & E. Ezcurra. 1997. Differentiationdiversity of Argentine cacti and its relationshipto
environmentalfactors.J. Veg. Sci. 8: 547-558.
Nassar, J. M., N. Ramirez & 0. Linares. 1997. Comparativepollination biology of Venezuelan colunmar cacti and the role of nectar-feedingbats in their sexual reproduction.Amer. J. Bot. 84:
918-927.
Negr6n-Ortiz, V. 1998. Reproductivebiology of a rarecactus, Opuntiaspinosissima(Cactaceae),in the
FloridaKeys: Why is seed set very low?. Sexual P1.Reprod. 11: 208-212.
Niering, W. A., R. H. Whittaker & C. H. Lowe. 1963. The saguaro:A populationin relationto environment. Science 142: 15-23.
Nobel, P. S. 1980. Morphology, nurse plant and minimum apical temperaturesfor young Carnegiea
gigantea. Bot. Gaz. 141: 188-191.
1988. Environmentalbiology of agaves and cacti. CambridgeUniversityPress. Cambridge,MA.
1989. Temperature,water availability,and nutrientlevels at various soil depths:Consequences
for shallow-rooteddesert succulents,includingnurse plant effects. Amer. J. Bot. 76: 1486-1489.
, G N. Gelier, S. C. Kee & A. D. Zimmerman. 1986. Temperaturesand thermaltolerancesfor
cacti exposed to high temperaturesnear the soil surface.P1.Cell Environ.9: 279-287.
Nolasco, H., F. Vega-Villasante & A. Diaz-Rondero. 1997. Seed germinationof Stenocereusthurberi
(Cactaceae)underdifferentsolar irradiationlevels. J. Arid Environm.36: 123-132.
Noy-Meir, I. 1973. Desert ecosystems: Environmentand producers.Annual Rev. Ecol. Syst. 4: 25-51.
. 1974. Desert ecosystems: highertrophiclevels. Annual Rev. Ecol. Syst. 5: 195-214.
9
199
1979/1980. Structureand function of desert ecosystems. Israel J. Bot. 28: 1-19.

Ortega-Baes, P. F. 2001. Demografiade la cactaceacolumnarEscontriachiotilla.M.S. thesis, Universidad
Nacional Aut6nomade Mexico, Mexico City.
Osborn, M. M., P. G Kevan & M. A. Lane. 1988. Pollination biology of Opuntiapolyacantha and
Opuntiaphaeacantha (Cactaceae)in southernColorado.PI. Syst. Evol. 159: 85-94.
Parker, K. C. 1987. Seedcrop characteristicsand minimum reproductive size of organ pipe cactus
(Stenocereusthurberi)in southernArizona. Madrofio34: 294-303.
. 1988a. Environmentalrelationshipsand vegetation associates of columnarcacti in the northern
SonoranDesert. Vegetatio78: 125-140.
. 1988b. Growthrates of Stenocereusthurberiand Lophocereusschottii in southernArizona.Bot.
Gaz. 149: 335-346.
.1989. Heightstructureandreproductivecharacteristicsof Senita,Lophocereusschottii(Cactaceae),
in southernArizona. Southw.Naturalist34: 392-401.
. 1991. Topography,substrate,and vegetationpatternsin the northernSonoranDesert.J. Biogeogr.
18: 151-163.
. 1993. Climatic effects on regenerationtrends for two columnarcacti in the northernSonoran
Desert.Ann. Assoc. Amer.Geogr. 83: 452-474.
& J. L. Hamrick. 1992. Genetic diversityand clonal structurein a columnarcactus,Lophocereus
schottii. Amer. J. Bot. 79: 86-96.
Pearl, R. 1928. The rate of living, being an account of some experimentalstudies on the biology of life
duration.Knopf, New York..
Pierson, E. A. & R. M. Turner. 1998. An 85-year study of saguaro(Carnegiea gigantea) demography.
Ecology 79: 2676-2693.
Pimienta, E., G Hernandez, A. Domingues & P. S. Nobel. 1998. Growth and development of the
arborescentcactus Stenocereus queretaroensisin a subtropicalsemiarid environment, including
effects of gibbberellicacid. Tree Physiol. 18: 59-64.
Reid, W., R. Lozano & R. Odom. 1983. Non-equilibriumpopulation structurein three Chihuahuan
Desert cacti. Southw.Naturalist28: 115-117.
Rinc6n, E., P. Huante & Y. Ramirez. 1993. Influence of vesicular-arbuscular
mycorrhizaeon biomass
productionby the cactus Pachycereuspecten-aboriginum.Mycorrhiza3: 79-81.
Rodriguez-Ortega, C. E. & E. Ezcurra. 2001. Distribuci6n espacial en el habitat de Mammillaria
pectiniferay M. camea en el Valle de ZapotitlanSalinas,Puebla,Mexico. Cact. Suc. Mex. 45: 4-14.
& M. Franco. 2001. La retenci6nde semillasen el generoMammillaria.Cact.Suc. Mex. 46: 63-67.
Rojas-Arechiga, M. & A. Batis. 2001. Las semillas de CactaceasJformanbancos en el suelo?. Cact. Suc.
Mex. 46: 76-82.
& C. Vazquez-Yanes. 2000. Cactus seed germination:A review. J. Arid Environm.44: 85-104.
A. Casas & C. Vazquez-Yanes. 2001. Seed germinationof wild and cultivated Stenocereus
stellatus (Cactaceae)from the Tehuacan-Cuicatlan
Valley, CentralMexico. J. Arid Environm.49:
279-287.
Rojas-Martinez, A., A. Valiente-Banuet, M. C. Arizmendi, A. Alcantara-Eguren & H. T. Arita. 1999.
Seasonal distributionof the long-nosed bat (Leptonycteriscurasoae) in North America: Does a
generalizedmigrationpatternreally exist?. J. Biogeogr. 26: 1065-1077.
Rosas-Barrera, M. L. & M. C. Mandujano. 2002. La diversidadde historias de vida de Cactaceas,
aproximaci6npor el triangulodemografico.Cact. Suc. Mex. 47: 33-41.
Ruedas, M., T. Valverde & S. C. Argulero. 2000. Respuesta germinativay crecimientotempranode
plantulasde Mammillariamagnimamma(Cactaceae)bajo diferentescondiciones ambientales.Bol.
Soc. Bot. Mexico 66: 25-35.
Ruiz, A., M. Santos, J. Cavalier & P. J. Soriano. 2000. Estudiofenol6gico de Cactaceasen el enclave
seco de la Tatacoa,Colombia.Biotropica32: 397-407.
Sahley, C. T. 1996. Bat and hummingbirdpollination of an autotetraploidcolumnar cactus, Werberbaurocereuswerberbaueri(Cactaceae).Amer. J. Bot. 83: 1329-1336.
Schlindwein, C. & D. Wittmann. 1997. Stamenmovements in flowers of Opuntia(Cactaceae)favour
oligolectic pollinators.PI. Syst. Evol. 204: 179-193.
Schmalzel, R. J., F. W. Reichenbacher & S. Rutman. 1995. Demographicstudyof the rareCoryphantha
robbinsorum(Cactaceae)in southeasternArizona. Madroflo42: 332-348.
200
Shreve, F. 1910. The rate of establishmentof the giant cactus. P1.World 10: 235-240.
. 1931. Physical conditions in sun and shade. Ecology 12: 96-104.
Silva, W. R. 1988. Ornitocoriaem Cereusperuvianus(Cactaceae)na Serrado Japi, Estadode Sao Paulo.
Revista Brasil. Biol. 48: 381-389.
Silvertown, J. W. 1987. Introductionto plant populationecology. Ed. 2. LongmanScientific & Technical, Essex, England.
& J. B. Wilson. 1994. Communitystructurein a desertperennialcommunity.Ecology 75: 409-417.
M. Franco, I. Pisanty & A. Mendoza. 1993. Comparativeplant demography:Relative importance of life-cycle componentsto the finite rate of increasein woody and herbaceousperennials.J.
Ecol. 81: 465-476.
Silvius, K. M. 1995. Avian consumers of card6n fruits (Stenocereusgriseus. Cactaceae)on Margarita
Island, Venezuela.Biotropica27: 96-105.
Spears, E. E., Jr. 1987. Island and mainlandpollinationecology of Centrosemavirginianumand Opuntia stricta. J. Ecol. 75: 351-362.
Steenbergh, W. F. & C. H. Lowe. 1969. Critical factors during the first years of life of the saguaro
(Cereusgiganteus) at SaguaroNational Monument,Arizona. Ecology 50: 823-834.
&
. 1977. Ecology of the saguaro:II, Reproduction,germination,establishment,growth,
and survivalof the young plant.Departmentof the Interior,NationalParkService, Washington,DC.
Suzain,H., G P. Nabhan & D. T. Patten. 1996. The importanceof Olneya tesota as a nurseplant in the
SonoranDesert. J. Veg. Sci. 7: 635-644.
Tinoco-Ojanguren, C. & F. Molina-Freaner. 2000. Flower orientationin Pachycereuspringlei. Canad.
J. Bot. 78: 1489-1494.
Tschapka, M., 0. von Helversen & W. Barthlott. 1999. Bat pollination of Weberocereustunilla, an
epiphyticrain forest cactus with functionalflagelliflory.P1.Biol. (Germany)1: 554-559.
Turner, R. M., S. M. Alcorn, G Olin & J. A. Booth. 1966. The influence of shade, soil, and water on
saguaroseedling establishment.Bot. Gaz. 127: 95-102.
Valiente-Banuet, A. & E. Ezcurra. 1991. Shade as a cause of the association between the cactus
Neobuxbaumiatetetzoand the nurseplantMimosa luisana in the TehuacanValley,Mexico. J. Ecol.
79: 961-971.
& H. Godinez-Alvarez. 2002. Populationand communityecology. Pp. 91-108 in P. S. Nobel
(ed.), Cacti:Biology and uses. University of CaliforniaPress, Berkeley.
, A. Bolongaro-Crevenna, 0. Briones, E. Ezcurra, M. Rosas, H. Nuinez,G Barnard & E. Vizquez. 1991a. Spatial relationshipsbetween cacti and nurse shrubs in a semi-aridenvironmentin
centralMexico. J. Veg. Sci. 2: 15-20.
, F. Vite & J. A. Zavala-Hurtado. 1991b. Interactionbetween the cactus Neobuxbaumiatetetzo
and the nurse shrubMimosa luisana. J. Veg. Sci 2: 11-14.
, P. Divila, R. J. Ortega, M. C. Arizmendi, J. L. Le6n, A. Breceda & J. Cancino. 1995.
Influencia de la evoluci6n de una pendiente de piedemonte en una vegetaci6n de cardonal de
Pachycereuspringlei en Baja CaliforniaSur, Mexico. Investigaci6nGeogr. Bol. 3: 101-113.
, M. C. Arizmendi, A. Rojas-Martinez & L. Dominguez-Canseco. 1996. Ecological relationships between columnarcacti and nectar-feedingbats in Mexico. J. Trop.Ecol. 12: 103-119.
, A. Rojas-Martinez, M. C. Arizmendi & P. Davila. 1997a. Pollinationbiology of two columnar
cacti (NeobuxbaumiamezcalaensisandNeobuxbaumiamacrocephala)in the TehuacanValley,central
Mexico. Amer. J. Bot. 84: 452-455.
' A. Casas, M. C. Arizmendi & P. Davila. 1997b. Pollination biology of two winterblooming giant columnar cacti in the TehuacanValley, central Mexico. J. Arid Environm. 37:
331-341.
, M. C. Arizmendi, A. Rojas-Martinez, A. Casas, H. Godinez-Alvarez, C. Silva & P. Davila.
2002. Biotic interactionsand populationdynamicsof columnarcacti. Pp. 225-240 in T. H. Fleming
& A. Valiente-Banuet(eds.), Columnarcacti and theirmutualists:Evolution,ecology, and conservation. Universityof Arizona Press, Tucson.
Valverde, T. & J. Silvertown. 1998. Spatial variation in the demographyof a forest understoryherb
(Primula vulgaris):Projectionmatrixmodels. J. Ecol. 86: 545-562.
, M. L. Trejo & S. Castillo. 1999. Patr6n de distribuci6n y abundancia de Mammillaria
magnimammaen la reservadel pedregalde San Angel, Mexico D. F. Cact. Suc. Mex. 44: 64-74.
201
S. Quijas, M. Lopez-Villavicencio & S. Castillo. In press. Populationdynamics of Mammillaria magnimammaHaworth.(Cactaceae)in a Mexican lava-field. P1.Ecol.
Weiss, J., A. Nerd & Y. Mizrahi. 1994. Flowering behavior and pollinationrequirementsin climbing
cacti with fruit crop potential.HortScience29: 1487-1492.
Yeaton, R. I. 1978. A cyclical relationshipbetween Larrea tridentataand Opuntia leptocaulis in the
northernChihuahuanDesert. J. Ecol. 66: 651-656.
Zavala-Hurtado, J. A. & A. Diaz-Solis. 1995. Repair,growth,age and reproductionin the giant columnar cactus Cephalocereuscolumna-trajani(Karwinskiex. Pfeiffer) Schumann(Cactaceae).J. Arid
Environm.31: 21-31.
XI. Appendix 1: Demographic information, endangerment, and geographical distribution of cac

Patternof distribution:1= random,2 = regular,3 = clumped; Recruitment:1 = under the canopy of tre
vegetation; Survivorship:1 = mortality is higher in the last size/age categories than in the first ones, 2 =
3 = mortality is higher in the first age/size categories than in later ones; Fecundity: 1 = fecu
2 = fecundity increases until a certain size/age and then remains constant, 3 = fecundity increases
then decreases; Size structure:1 = numberof individuals decreases with size/age, 2 = number of
3 = numberof individuals varies with size/age; Population dynamics: 1 = species analyzed with ma
in appendix I; IUCN: e = endangered,v = vulnerable, r = rare, i = indeterminate;Geographica
S = South America, * = endemic to one country of the geographical r
Species
Globose cacti
Ancistrocactus tobuschii
Ariocarpus trigonus
Coryphantharobbinsorum
Mammillariacrucigera
M gaumeri
Endangered
species
Patternof Recruit- SurvivorSize
Population
distribution ment
Fecundity structure dynamics CITES IUCN
ship
1
1
3
2/3
M magnimamma
M microcarpa
Neolloydia pseudopectinata
3
3
1
2
Barrelcacti
Copiapoa cinerea
Echinocactus horizonthalonius
Echinocereus engelmanni
E. triglochidiatus
I
3
1/3
3
v
e
r
1
3
v
v
e
i
Echinomastus erectrocentrus
Ferocactus acanthodes
Cephalocereus columnatrajani
Escontria chiotilla
Lophocereusschottii
Neobuxbaumiamacrocephala
N. tetetzo
Pachycereuspringlei
Stenocereusthurberi
Trichocereuspasacana
Opuntioidcacti
Opuntiaechios
0. rastrera
F cylindraceus
F histrix
F wislizeni
Columnarcatci
Carnegiea gigantea
3
1
3
3
1
1
1
3
3
3
1
I
3
1
3
1
1/3

Demographic Trends in The Cactaceae PDF

Hochgeladen von

Dokumentinformationen

Originalbeschreibung:

Originaltitel

Copyright

Verfügbare Formate

Dieses Dokument teilen

Dokument teilen oder einbetten

Freigabeoptionen

Stufen Sie dieses Dokument als nützlich ein?

Sind diese Inhalte unangemessen?

Copyright:

Verfügbare Formate

Demographic Trends in The Cactaceae PDF

Hochgeladen von

Copyright:

Verfügbare Formate

Demographic Trends in the Cactaceae

Author(s): Hctor Godnez-lvarez, Teresa Valverde, Pablo Ortega-Baes

The Botanical Review 69(2): 173-203

Demographic Trends in the Cactaceae

Issued 16 December 2003

THE BOTANICAL REVIEW

aparchonadade los recursosen los ambientesaltamenteheterogeneosquehabitan.La asociaci6n

THE BOTANICAL REVIEW

minationandearlygrowth(Rojas-Arechiga& Vazquez-Yanes,2000; Ruedaset al., 2000; Flores

III. Distribution and Abundance Patterns

THE BOTANICAL REVIEW

Contrastingly,Haman(2001) reportsthatEl Nino events have an adverseeffect on two species

IV. Seed Germination and Seedling Establishment

Association between nurse plants and different cacti species

Life-cycle stage Mechanism

Increased soil nutrients

Life-cycle stage Mechanism

Seed dispersal site?

Despite the fact thatnurseplantsmay offer cacti some protectionduringthe initiallife-cycle

V. Growth and Reproduction

DEMOGRAPHYIN THE CACTACEAE

Johnson et al., 1992

Schmalzel et al., 1995

Plant size measuredas the numberof branches.

Lophocereus schottii, Neobuxbaumia macrocephala, N. tetetzo, and Stenocereus thurberi

gigantea, Echinomastuserectrocentrus,Lophocereusschottii,Mammillariacrucigera,M mag-

THE BOTANICAL REVIEW

THE BOTANICAL REVIEW

Nassar et al., 1997

McFarlandet al., 1989

Schlindwein & Wittmann,1997

Osbom et al., 1988

Steenbergh& Lowe, 1977

Nassar et al., 1997

Osbom et al., 1988

Schlindwein & Wittmann,1997

DEMOGRAPHY IN THE CACTACEAE

Fig. 2. Populationsize structures.a. The barrelcactus Echinocactushorizonthalonius.b. The globose

VII. Population Dynamics

Summary of population dynamics data for 17 cactus populations of eight specie

Per demographic proces

THE BOTANICAL REVIEW

Fig. 3 (above). Triangularordinationof elasticityvalues for the 17 matricesreportedin TableIV. The

DEMOGRAPHY IN THE CACTACEAE

THE BOTANICAL REVIEW

In additionto evaluatingthe contributionof particulardemographicprocesses, as proposed

. 1996b. Environmentaldeterminantsof floweringdate in the columnarcactus Carnegieagigantea

DEMOGRAPHY IN THE CACTACEAE

Flores, F. J. L. & R. I. Yeaton. 2000. La importanciade la competencia en la organizaci6n de las

Larmuth,J. & H. J. Harvey. 1978.Aspectsof the occurrenceof desertplants.J.AridEnvironm.1: 129-133.

DEMOGRAPHYIN THE CACTACEAE

1979/1980. Structureand function of desert ecosystems. Israel J. Bot. 28: 1-19.

THE BOTANICAL REVIEW

DEMOGRAPHYIN THE CACTACEAE

XI. Appendix 1: Demographic information, endangerment, and geographical distribution of cac

Das könnte Ihnen auch gefallen